Abstract
Drought stress profoundly affects plant growth and survival, but comparisons of integrated adaptive strategies across multiple tree species remain unclear. In this study, seedlings of Elaeagnus angustifolia (E. angustifolia), Populus euphratica (P. euphratica) and Xanthoceras sorbifolium (X. sorbifolium) were subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments. Leaf microanatomical traits, non-structural carbohydrates (NSCs), stoichiometric elements, biomass allocation, and key stress indicators were measured. The results showed that P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%); their antioxidant enzymes were activated only under mild-to-moderate stress and declined under severe stress. E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass, continuously elevated activities of superoxide dismutase (SOD) and peroxidase (POD) as well as osmoregulatory substances (soluble protein SP, proline Pro), and showed the lowest malondialdehyde (MDA) content; their leaf carbon (C), nitrogen (N), and phosphorus (P) contents decreased the least, and their stoichiometric ratios remained stable. In contrast, X. sorbifolium seedlings progressively reduced leaf thickness and vascular area, depleted NSC reserves, exhibited unstable antioxidant responses, showed a significant decrease in Pro under severe drought, accumulated the highest MDA, and had the lowest N/P ratio, indicating the strongest nitrogen limitation. These results demonstrate that E. angustifolia combines structural plasticity, efficient nutrient use, robust osmotic adjustment, and sustained antioxidant capacity, conferring the strongest drought tolerance; P. euphratica* shows moderate tolerance through transient structural and carbon investment but suffers under extreme drought; X. sorbifolium has the weakest drought tolerance.
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Claims 20
Claim · model-written summary
Populus euphratica thickened leaves and vascular tissues under mild and moderate drought (T1-T2) compared with well-watered controls.
- Taxon
- P. euphratica
Populus euphraticae1 - Treatment
- T1–T2e1
- Outcome
- leaves and vascular tissuese1
- Direction
- thickenede1
- Comparator
- well-watered (CK)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”
e1“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”
Claim · model-written summary
Populus euphratica accumulated soluble sugars and starch under mild and moderate drought (T1-T2) compared with well-watered controls.
- Taxon
- P. euphratica
Populus euphraticae1 - Treatment
- T1–T2e1
- Outcome
- soluble sugars (SSs) and starch (ST)e1
- Direction
- accumulatede1
- Comparator
- well-watered (CK)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”
e1“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”
Claim · model-written summary
Populus euphratica increased root biomass by 26.0% under severe drought (T3), prioritizing root investment.
- Taxon
- P. euphratica
Populus euphraticae1 - Treatment
- T3e1
- Outcome
- root biomasse1
- Direction
- increaseinferrede1
- Value
- +26.0%e1
- Comparator
- well-watered (CK)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”
e1“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”
Claim · model-written summary
Populus euphratica antioxidant enzymes were activated under mild-to-moderate stress.
- Taxon
- P. euphratica
Populus euphraticae0 - Treatment
- mild-to-moderate stresse1
- Outcome
- antioxidant enzymese1
- Direction
- activatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”
e1“their antioxidant enzymes were activated only under mild-to-moderate stress and declined under severe stress”
Claim · model-written summary
Populus euphratica antioxidant enzymes declined under severe stress.
- Taxon
- P. euphratica
Populus euphraticae0 - Treatment
- severe stresse1
- Outcome
- antioxidant enzymese1
- Direction
- declinede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“P. euphratica seedlings thickened leaves and vascular tissues and accumulated soluble sugars (SSs) and starch (ST) under T1–T2, but under T3, they prioritized root investment (root biomass +26.0%)”
e1“their antioxidant enzymes were activated only under mild-to-moderate stress and declined under severe stress”
Claim · model-written summary
Elaeagnus angustifolia increased root biomass by 97.2% under severe drought (T3), with moderate leaf structural thickening.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae1 - Treatment
- T3e1
- Outcome
- root biomasse1
- Direction
- increasede1
- Value
- +97.2%e1
- Qualifier
- moderate leaf structural thickeninge1
- Comparator
- well-watered (CK)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“subjected to well-watered (CK), mild (T1), moderate (T2), and severe (T3) drought treatments”
e1“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
Claim · model-written summary
Elaeagnus angustifolia maintained stem biomass under severe drought (T3).
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Treatment
- T3e0
- Outcome
- stem biomasse0
- Direction
- no_changeinferrede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
Claim · model-written summary
Elaeagnus angustifolia showed continuously elevated superoxide dismutase and peroxidase activities.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Outcome
- activities of superoxide dismutase (SOD) and peroxidase (POD)e1
- Direction
- elevatede1
- Qualifier
- continuouslye1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
e1“continuously elevated activities of superoxide dismutase (SOD) and peroxidase (POD)”
Claim · model-written summary
Elaeagnus angustifolia showed continuously elevated soluble protein and proline.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Outcome
- soluble protein SP, proline Proe2
- Direction
- elevatede1
- Qualifier
- continuouslye1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
e1“continuously elevated activities of superoxide dismutase (SOD) and peroxidase (POD)”
e2“as well as osmoregulatory substances (soluble protein SP, proline Pro)”
Claim · model-written summary
Elaeagnus angustifolia had the lowest malondialdehyde content.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Outcome
- malondialdehyde (MDA) contente1
- Direction
- loweste1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
e1“showed the lowest malondialdehyde (MDA) content”
Claim · model-written summary
Elaeagnus angustifolia leaf carbon, nitrogen and phosphorus contents decreased the least under drought.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Outcome
- leaf carbon (C), nitrogen (N), and phosphorus (P) contentse1
- Direction
- decreased the leaste1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
e1“their leaf carbon (C), nitrogen (N), and phosphorus (P) contents decreased the least”
Claim · model-written summary
Elaeagnus angustifolia stoichiometric ratios remained stable under drought.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Outcome
- stoichiometric ratiose1
- Direction
- remained stablee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia seedlings exhibited moderate leaf structural thickening, sharply increased root biomass (+97.2% under T3) while maintaining stem biomass”
e1“their stoichiometric ratios remained stable”
Claim · model-written summary
Xanthoceras sorbifolium progressively reduced leaf thickness and vascular area under drought.
- Taxon
- X. sorbifoliumXanthoceras sorbifoliume0
- Outcome
- leaf thickness and vascular areae0
- Direction
- progressively reducede0
Verbatim
e0“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”
Claim · model-written summary
Xanthoceras sorbifolium depleted nonstructural carbohydrate reserves under drought.
- Taxon
- X. sorbifoliumXanthoceras sorbifoliume0
- Outcome
- NSC reservese1
- Direction
- depletede1
Verbatim
e0“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”
e1“depleted NSC reserves”
Claim · model-written summary
Xanthoceras sorbifolium showed a significant decrease in proline under severe drought.
- Taxon
- X. sorbifoliumXanthoceras sorbifoliume0
- Treatment
- severe droughte1
- Outcome
- Proe1
- Direction
- decreasee1
Not reportedComparator
Verbatim
e0“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”
e1“showed a significant decrease in Pro under severe drought”
Claim · model-written summary
Xanthoceras sorbifolium accumulated the highest malondialdehyde content.
- Taxon
- X. sorbifoliumXanthoceras sorbifoliume0
- Outcome
- MDAe1
- Direction
- higheste1
Verbatim
e0“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”
e1“accumulated the highest MDA”
Claim · model-written summary
Xanthoceras sorbifolium had the lowest N/P ratio, indicating the strongest nitrogen limitation.
- Taxon
- X. sorbifoliumXanthoceras sorbifoliume0
- Outcome
- N/P ratioe1
- Direction
- loweste1
- Qualifier
- indicating the strongest nitrogen limitatione1
Verbatim
e0“X. sorbifolium seedlings progressively reduced leaf thickness and vascular area”
e1“had the lowest N/P ratio, indicating the strongest nitrogen limitation”
Claim · model-written summary
Elaeagnus angustifolia has the strongest drought tolerance of the three species.
- Taxon
- E. angustifolia
Elaeagnus angustifoliae0 - Outcome
- drought tolerancee0
- Direction
- strongeste0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“E. angustifolia combines structural plasticity, efficient nutrient use, robust osmotic adjustment, and sustained antioxidant capacity, conferring the strongest drought tolerance”
Claim · model-written summary
Populus euphratica shows moderate drought tolerance but suffers under extreme drought.
- Taxon
- P. euphratica
Populus euphraticae0 - Outcome
- tolerancee0
- Direction
- moderatee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“P. euphratica* shows moderate tolerance through transient structural and carbon investment but suffers under extreme drought”
Claim · model-written summary
Xanthoceras sorbifolium has the weakest drought tolerance of the three species.
- Taxon
- X. sorbifoliumXanthoceras sorbifoliume0
- Outcome
- drought tolerancee0
- Direction
- weakeste0
Verbatim
e0“X. sorbifolium has the weakest drought tolerance”